REVISION OF FEVILLEA (CUCURBITACEAE: ZANONIEAE)

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1 REVISION OF FEVILLEA (CUCURBITACEAE: ZANONIEAE) George L. Robinson Richard P. Wunderlin USDA APHIS, Plant Protection and Quarantine Department of Biology, SCA 110 Port Everglades University of South Florida 1800 Eller Drive, Suite 414 Tampa, Florida , U.S.A. Fort Lauderdale, Florida 33316, U.S.A. ABSTRACT A revision of the neotropical genus Fevillea with keys, descriptions, and distribution is provided. Two subgenera are recognized: subgenus Fevillea with six species and subgenus Anisosperma with one species. Fevillea bahiensis G. Robinson & Wunderlin from Brazil is described as new and subgenus Anisosperma (Silva Manso) G. Robinson & Wunderlin is proposed. RESUMEN Se aporta una revisión del género neotropical Fevillea con claves, descripciones, y distribución. Se reconocen dos subgéneros: subgénero Fevillea con seis especies y subgénero Anisosperma con una especie. Se describe como nueva Fevillea bahiensis G. Robinson & Wunderlin de Brasil y se propone el subgénero Anisosperma (Silva Manso) G. Robinson & Wunderlin. Fevillea (Cucurbitaceae: Zanonieae), a neotropical genus of seven species, is characterized by leaves with glands, calyx with glandular squamellae (except F. passiflora), petals with a medial adaxial flap-like appendage or ridge, staminate flowers with five bilocular anthers, and a globose, usually indehiscent, large-seeded fruit. TAXONOMIC HISTORY Fevillea was established by Linnaeus (1753) in honor of Louis Éconches Feuillée, ( ), a French clergyman, explorer, astronomer, and botanist. Linnaeus recognized two species, F. trilobata and F. cordifolia. Adanson (1763) published the pre-linnaean name Nhandiroba, of Marcgrave (Piso & Marcgrave 1648), but placed Linnaeus s name Fevillea in synonymy, thereby making Nhandiroba illegitimate. No species were listed. The name Nhandiroba remained unused until resurrected by Kuntze ( ). The first comprehensive treatment of Fevillea since Linnaeus (1753) was that of Seringe (1828) who recognized four species. In addition to F. cordifolia and F. trilobata, Seringe recognized F. punctata (L.) Poir. [=Trichosanthes sp.] and F. javilla Kunth [=F. cordifolia]. Silva Manso (1836) established the monotypic Hypanthera with H. guapeva [=Fevillea trilobata] and Anisosperma with A. passiflora (Vell.) Silva Manso [=Fevillea passiflora Vell.]. Both monotypic genera were accepted by most subsequent workers until recently. SIDA 21(4):

2 1972 BRIT.ORG/SIDA 21(4) Roemer (1846) recognized Hypanthera and Fevillea. He placed ten species in Fevillea, incorporating some taxa now placed in Trichosanthes L., Pteropepon Cogn., and Sicydium Schltdl. Fevillea passiflora was also recognized, but without reference to Silva Manso s placement of the species in Anisosperma. Roemer erected two sections: section Fevillea with nine species and section Javilla with only F. javilla Kunth [=F. cordifolia L.]. Cogniaux (1878) recognized three species of Fevillea for Brazil: F. trilobata, F. albiflora Cogn., and F. deltoidea Cogn. (the latter now in Pteropepon). In addition, he recognized the monotypic genus Anisosperma and expanded the generic limits of the previously Old World genus Alsomitra (Blume) M. Roem. to include two new neotropical species: A. brasiliensis [=Siolmatra brasiliensis (Cogn.) Baill.] and A. pedatifolia [=Fevillea pedatifolia (Cogn.) C. Jeffrey]. In a more comprehensive treatment, Cogniaux (1881) expanded Fevillea to six species, maintaining Anisosperma and Alsomitra. The genus Siolmatra, a segregate of Alsomitra created by Baillon (1885) to acommodate Alsomitra brasiliensis Cogn. [=Siolmatra brasiliensis (Cogn.) Baill.], was accepted by Cogniaux (1893) who described a second species (S. paraguayensis). Three additional species were later added by Cogniaux (1916), one new (S. amazonica) and two transferred from Alsomitra (S. pedatifolia (Cogn.) Cogn. and S. peruviana (Huber) Cogn.). Harms (1926) added a sixth species (S. pentaphylla), later (Harms 1933) a seventh (S. simplicifolia), and Standley (1937) an eighth (S. mexiae). Jeffrey (1962b), noting the incongruent mixture of taxa in Siolmatra, recircumscribed the genus and transferred four species to Fevillea. In reviewing the New World taxa of the Cucurbitaceae, Jeffrey (1978) listed nine species in Fevillea, including F. passiflora, considering Anisosperma congeneric with Fevillea. FLORAL AND FRUIT MORPHOLOGY The staminate inflorescence consists of numerous, small, pentamerous flowers that are paniculate in subgenus Fevillea or fasciculate to subumbelliform in subgenus Anisosperma. In subgenus Fevillea, the midrib of the adaxial surface of the sepals is fused with the lower margins of the petals. At or above this point of fusion extends a small glandular protuberance of uncertain ontogenic origin which is here referred to as a glandular calycine squamella. The exudate from the squamellae is clear and remains visible on most herbarium specimens. Each petal has a median, adaxial, uncinate appendage or slightly raised glandular midrib which is adnate with the base of the stamen filament. In subgenus Anisosperma, the sepals and petals are united at their base and lack squamellae. Instead, the petals have a median, adaxial, glandular ridge. An articulation occurs between the filiform hypanthium and the pedicel. The indumentum on the staminate flowers is quite variable in most species and

3 ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1973 is similar (when present) on both the hypanthium and pedicel. However, F. trilobata has a stipitate-glandular pubescence on the hypanthium above the articulation in rather sharp contrast to the non-glandular trichomes on the pedicel below. The presence of five free, bilocular anthers in Fevillea is unique for the family and is considered plesiomorphic. The general trend within the Cucurbitaceae is the reduction of stamen number from five to three or to two and the reduction in locule number from two to one. The pistillate flowers are rarely collected and are thus imperfectly known or unknown for some species. We have seen them in only four of the seven species. Those of F. pergamentacea and F. passiflora are known to us only through the literature while those of F. moorei are unknown. The petals of the pistillate flowers have a slightly raised median appendage extending from the base to the center, reminiscent of those of the staminate flowers. Two small glands occur at the base of the petal on either side of this median appendage. A large, subrotund, flattened, glandular protuberance extends from the base of the calyx lobe between each petal. These protuberances may represent staminodes. Cogniaux (1878, 1881, 1916) reported these structures, counting 20 small glands at the base of the petals. This probably included the sum total of glands, ridges, and protuberances. The fruits of subgenus Fevillea are large, subglobose, mottled green or brown, and gourd-like. The size ranges from 8 to 16 cm in length and from 7 to 13 cm in diameter. An individual plant may produce as many as 50 to 100 fruits at a time (Gentry & Wettach 1986). The fleshy rind of the fruit is zonate above the middle with the hypanthium lip scar (ovary partly inferior). The fruits are typically indehiscent, but reportedly sometimes dehiscent along the hypanthium lip scar in F. pedatifolia (A. Gentry, pers. comm.). In contrast, the fruits of subgenus Anisosperma are ovoid or oblong, subtrigonous, short-apiculate at the apex, and not zonate above the middle (ovary fully inferior). The seeds (up to 15 per fruit) vary from 3 to 6 cm in diameter and weigh 3 to 9 g when dry. They are among the largest in the Cucurbitaceae and are comparable in size only to those of the paleotropical genera Telfairia (Cucurbitoideae: Joliffieae) and Hodgsonia (Cucurbitoideae: Trichosantheae). The seed coat consists of three layers. The innermost layer surrounding the cotyledon is spongy and aeriferous. This layer is enclosed in a thin, hard, woody layer. The outermost layer is thin, smooth, and of a corky texture which tends to obscure the margin of the woody layer below but usually does not persist. DISTRIBUTION AND HABITAT Fevillea cordifolia has the widest distribution, ranging from southern Mexico, east into the Caribbean to Puerto Rico, south through Central America and into South America to northern Argentina. Dieterle (1976) notes that it is cultivated

4 1974 BRIT.ORG/SIDA 21(4) in some or all Central American countries and is found in most Central American markets, especially in Guatemala. Three species (F. trilobata, F. bahiensis, and F. passiflora) are endemic to eastern Brazil and F. pedatifolia and F. pergamentacea occur in Ecuador, Peru, and Bolivia, with F. pedatifolia extending into adjacent Amazonian Brazil. Fevillea moorei is known only from the type material cultivated in England and is probably from Guyana or Amazonian Brazil. Fevillea typically occurs along river banks, along the edge of tropical primary or secondary forests, and along the edge of seasonally inundated riverine forests, occasionally climbing to heights of 35 m in forest canopy openings. It also is found in forest clearings and along roadsides. It occurs at elevations from near sea level to about 500 m, less commonly up to 1,700 meters. Fruits and seeds of Fevillea are quite buoyant and thus apparently are well suited to dispersal in fresh water. Gentry and Wettach (1986) report that at least one species (F. cordifolia or F. pedatifolia) of Amazonian Peru occurs in seasonally inundated forests, a habitat in which water dispersal is prevalent. Seed drift materials of Fevillea cordifolia have been found within the Caribbean basin well outside the species natural range. Gunn and Dennis (1976), Morton (1981), and the senior author have identified seeds of F. cordifolia collected from beaches of southern Florida. Guppy (1917) reported materials found along beaches of the Turks Islands, Tobago, and Grenada. Guppy (1917) and Gunn and Dennis (1976) found that seeds of F. cordifolia germinated in fresh water while afloat after the disintegration of the fruit wall, but were generally rendered non-viable in salt water. Although F. cordifolia is sometimes listed as an estuarine plant, the seeds are probably not capable of over-sea transport for any distance. However, dispersal by seed drift via salt water can not be disregarded. Guppy (1917) estimated that 5% of the Fevillea cordifolia drift seeds reaching the Turks Islands were viable while Gunn and Dennis (1976) found 20% of the undamaged drift seeds on Florida beaches were viable. The high salinity of the beach is probably lethal, thus preventing colonization. ECONOMIC IMPORTANCE The high seed oil content of Fevillea trilobata was recognized centuries ago by indigenous Brazilians whose use of it was first documented by Marcgrave (Piso & Marcgrave 1648). Fevillea cordifolia is similarly well known in the ethnobotanical literature (cf. Gentry & Wettach 1986). It has been used as a purgative, reputed antidote for many kinds of poisoning, and as a treatment for numerous diseases. In Jamaica it is called antidote caccoon or antidote vine (Adams 1972; Gunn & Dennis 1976; Morton 1981). Lindley and Moore (1870) first reported the use of Fevillea seeds by Peruvians as candles. Gentry and Wettach (1986) report that abiria (Fevillea pedatifolia) is used as candles by the Campa Indians of the Pichis Valley of Peru. Fevillea seeds may have potential as an edible or fuel oil source. Calculated

5 ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1975 on a weight per fruit basis, the seed oil content of Fevillea is higher than that of any other dicotyledon (Gentry & Wettach 1986). Preliminary analysis of oils extracted from the seeds of two Peruvian species (F. cordifolia and F. pedatifolia) by Gentry and Wettach show the oil to be simple triglycerides, slightly heavier than those of refined cottonseed oil. All species were rich in the saturated lowweight fatty acids, palmitic (21 60%) and stearic (10 42%), and the unsaturated oleic (17 17%) and linoleic (6 7%). When compared with previous results from the Brazilian F. trilobata (Tulloch & Bergter 1979), Gentry and Wettach suggest that the Peruvian species with 60 70% low-weight, saturated fatty acids would be a good sources of fuel oil while the Brazilian species with 57% unsaturated fatty acids would be a good source of polyunsaturated edible oils. However, the relatively high concentration of high molecular weight fatty acids in F. trilobata, probably correlated with its purgative properties, may reduce its value as an edible oil. The high percent of stearic acid in F. cordifolia might also suggest its use in the candle industry where this chemical is used to harden waxes and in the rubber industry where used as an extender. Preliminary laboratory analysis of F. cordifolia at the University of South Florida gave highly variable results thought to be related to the different ages of the seeds tested, further complicating the use of Fevillea seeds as a potential commercial oil source (unpublished data). Although Fevillea as an oil source is documented in the literature, to date it remains a genus of little or no economic importance. INFRAGENERIC RELATIONSHIPS Jeffrey (1962a) recognized two genera within the subtribe Fevilleinae, Fevillea and Anisosperma. In our treatment, Anisosperma is reduced to a subgenus of Fevillea. Subgenus Anisosperma differs from subgenus Fevillea by the shape of the corolla lobes, the character of the median adaxial glandular ridge of the staminate petals, the congested staminate inflorescence, the lack of glandular calycine squamellae on the staminate flowers, and the fruit shape. However, with the presence of the foliar glands, the median adaxial glandular ridge on the petals, and overall similarity in fruit and seed morphology, the single species of subgenus Anisosperma is easily accommodated in Fevillea. Within subgenus Fevillea, three species groups can be distinguished on the basis of foliar gland characters. The first group consists of F. pergamentacea and F. pedatifolia which have conspicuous petiolar glands and inconspicuous laminar glands. The others are characterized by having laminar glands only. Of these, F. cordifolia and F. trilobata have glands terminating the veins on the lamina and lack basal laminar glands while F. moorei and F. bahiensis both have basal laminar glands only. SYSTEMATIC TREATMENT Fevillea L., Sp. Pl Nhandiroba Adan., Fam. Pl. 2: , nom. illegit.

6 1976 BRIT.ORG/SIDA 21(4) TYPE: Fevillea trilobata L. Lectotype designated by M.L. Green (in Sprague et al., Nom. Prop. Brit. Bot ). This lectotypification replaces the lectotype of Fevillea cordifolia L. of Britton and P. Wilson. (Sci. Surv. Porto Rico 6: ) under Art. 10.5b of the St. Louis Code. Anisosperma Silva Manso, Enum. Subst. Braz TYPE: Anisosperma passiflora (Vell.) Silva Manso [=Fevillea passiflora Vell.]. Hypanthera Silva Manso, Enum. Subst. Braz TYPE: Hypanthera guapeva Silva Manso [=Fevillea trilobata L.]. Dioecious vines or lianas; stems sulcate; tendrils axillary, sulcate, distally 2-fid, coiling both above and below the bifurcation. Leaves alternate, petiolate, the blade unlobed, or palmately 3- to 7-lobed, or 3- to 5-foliolate, with glands on the leaf margins terminating the primary lateral veins and/or 2 glands at the blade base or on the petiole, the petiole canaliculate, sometimes bearing 2 glands at or above the middle. Staminate inflorescences paniculate or subumbelliform, many-flowered, bracteate; flowers short-pedicellate; hypanthium pedicelloid; calyx lobes 5, fused to the petals above, not completely enclosing the petals in bud, with a glandular calycine squamella on each calyx lobe at or near the point of fusion with the petals (except in F. passiflora); petals 5, fused to the sepals below, the lobes each with a median, adaxial, uncinate, flap-like appendage or a slightly raised ridge or (in F. passiflora) with a thick, glandular ridge; stamens 5, equal, free, inserted near the center of the flower, the anthers bilocular, extrorse, dehiscing longitudinally, the connective with an adaxial glandular protuberance or projection; pollen prolate, µ in length, tricolporate, coarsely striate. Pistillate flowers solitary or in pairs; hypanthium deeply cupular; sepals and petals as in the staminate flowers or sometimes the petals differing in shape; ovary partly inferior, 3-locular, the styles 3, free, outwardly curved, the stigmas reniform, capitate, the ovules pendulous, usually 4 in each locule. Fruit globose, gourd-like with a thick, fleshy rind, zonate above the middle with the hypanthium lip scar or non-zonate, indehiscent or rarely circumscissile dehiscent along the hypanthium lip scar; seeds large, orbicular, somewhat compressed, the seed coat consisting of a thick, spongy, aeriferous inner layer surrounded by a thin woody layer, and an outer, usually non-persistent layer, the lateral surface smooth or striate-verrucose, the outer edges smooth or tuberculate, the inner kernel disk-like, oily. Two subgenera are distinguished as follows: 1. Corolla lobes of the staminate flowers suborbicular, the base cuneate, with a median adaxial uncinate appendage or sharply defined ridge; glandular calycine squamellae present between the petals and the calyx lobes; staminate flowers in spreading panicles; fruit subglobose, zonate above the middle, the apex rounded subg. Fevillea 1. Corolla lobes of staminate flowers oblong-hastate, the base with a median adaxial glandular ridge broadening downward; glandular calycine squamellae absent; staminate flowers in congested panicles or subumbelliform; fruit ovoid to oblong, not zonate, the apex short-apiculate subg. Anisosperma

7 ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1977 Fevillea subgenus Fevillea Fevillea section Javilla M. Roem., Fam. Nat. Syn. Monogr. 2: TYPE: Fevillea javilla Kunth [=Fevillea cordifolia L.]. Staminate inflorescences paniculate; glandular calycine squamellae present; corolla lobes suborbicular with a median adaxial uncinate appendage or slightly raised ridge. Fruit subglobose, zonate above the middle, the apex rounded. 1. Leaves pedately 3- to 5-foliolate F. pedatifolia 1. Leaves lobed or unlobed, but not pedately foliolate. 2. Leaf blade without glands at the base or on the petiole, with marginal glands terminating the veins. 3. Leaf blade with angled or rarely with rounded lobes, the marginal glands inconspicuous; staminate flowers with the hypanthium densely stipitateglandular pubescent, the pedicel with non-glandular trichomes F. trilobata 3. Leaf blade unlobed or occasionally with rounded lobes, the marginal glands conspicuous; staminate flowers with the hypanthium and pedicel variously pubescent but not as above F. cordifolia 2. Leaf blade with glands at the base or on the petiole, with or without marginal glands terminating the leaf veins. 4. Leaves with glands only at the blade base, without marginal glands terminating the leaf veins of the blade. 5. Staminate flowers 3 5 mm wide; leaves drying reddish-brown, the blade with a conspicuous uncinate-ciliate margin F. bahiensis 5. Staminate flowers mm wide; leaves drying green, the blade with a smooth margin F. moorei 4. Leaves with glands either at the blade base or on the petiole, also with marginal glands terminating the veins of the blade. 6. Leaves with glands at the blade base F. pergamentacea 6. Leaves with glands on the petiole F. pedatifolia Fevillea bahiensis G. Robinson & Wunderlin, sp. nov. (Fig. 1). TYPE: BRAZIL. BAHIA: 8 km to the N of Ubaitaba on BR 101, 16 Jun 1972, dos Santos 2307 (HOLOTYPE: CEPEC; ISOTYPE: K). Species haec a Fevillea moorei Hook. f. differt floribus staminatis minoribus, foliis in siccitate badiis usque atrobrunneis marginibus uncinato-ciliatibus. Vine or liana; stem glabrous to lightly appressed golden brown-pubescent; tendrils glabrous to lightly pubescent. Leaves unlobed, the blade ovate, (2.5 ) (12) cm long, (3.5 )5.5 9 cm wide, membranaceous, drying dark brown to reddish brown, 5-nerved, the apex acuminate, the base cordate to truncate, the margin entire, with two irregularly shaped glands at the base near the petiole, the upper and lower surfaces glabrous or with scattered, appressed, golden brown trichomes, these usually denser along the leaf veins, the margin uncinate-ciliate, the petiole (1.5 )3.5 5 cm long, glabrous to lightly pubescent. Staminate flowers in a paniculate inflorescence on reduced subterminal, lateral branches, the branches subtended by a reduced leaf; pedicel mm long, glabrous or sparsely pubescent; bracts linear, ca. 1 mm long; hypanthium mm long, lightly golden brown-pubescent; calyx shallowly cupular, the lobes

8 1978 BRIT.ORG/SIDA 21(4) FIG. 1. Fevillea bahiensis. A. Habit, staminate inflorescence (dos Santos 2307). B. Staminate flower (dos Santos 2307). up to 1.5 mm long, 1 mm wide, glabrous to sparsely pubescent abaxially, the margin entire, the apex rounded, with scattered stipitate glands, with a glandular calycine squamella at or near the point of fusion with the petals; petals suborbicular, mm long, ca. 1.5 mm wide, cream-colored, the margin entire, the median adaxial ridge slightly raised; stamens cm long, the anthers ca mm long, slightly longer than wide, the filaments ca. 0.5 mm long. Pistillate flowers solitary; calyx deeply cupular, the lobes ca. 1.5 mm long, ca. 1 mm wide, dark brown, fleshy; petals strap-shaped, ca. 2 mm long, ca. 1.5 mm wide, cream-colored. Immature fruit subglobose, 2 4 cm long and wide, the surface smooth; mature fruit not seen. Distribution and ecology. Endemic to Brazil in southern Bahia.

9 ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1979 Additional specimens examined. BRAZIL. Bahia: Nova Esperança, São Lourenço, 32 km W of Wenceslau Guimarães, 26 Jul 2001, Mattos-Silva et al (NY); Ramal da Torre da Embratel, entrance 15 km from the Ubaitabe/Itacaré Road (BR 654), 5.8 km from the entrance, 6 Jun 1978, Mori & dos Santos (CEPEC, K, NY); km 3, Uruçuca-Taboquinha highway, 19 Jun 1972, dos Santos 2316 (CEPEC); access road to Torre da Embratel, entrance to the right of road to Ubaitabe/Itacaré (BR 654), 24 Sep 1977, dos Santos 3130 (CEPEC, K); Almadina Mata da Serra Pancadinho, 10 Mar 1971, Pinheiro 1076 (CEPEC, K). Fevillea bahiensis is most similar and probably most closely related to Fevillea moorei of Guyana Amazonian Brazil. Both species have glands only at the base of the leaf blade. Fevillea bahiensis differs by having smaller staminate flowers (petals mm long vs. ca. 1 cm long in F. moorei) and the leaves drying a dark reddish brown and with conspicuous uncinate-ciliate margins. Fevillea cordifolia L., Sp. Pl (Fig. 2). Fevillea scandens L., Sp. Pl. ed , nom. illegit. Nhandiroba scandens Descourt., Fl. Méd. Antilles, index 66 [t. 198]. 1829, nom. illegit. Nhandiroba cordifolia (L.) Kuntze, Revis. Gen. Pl. 1: Fevillea cordifolia L. var. typica Stehlé, M. Stehlé, & Quentin, Fl. Guadeloupe 2(3): , nom. inadmiss. TYPE: Habitat in America Calidiore (LECTOTYPE: Nhandiroba, Plumier, Pl. Amer. 20, t ). Lectotypified by C. Jeffrey, (in C.E. Jarvis et al., eds. Regnum Veg. 127: ). Fevillea hederacea Poir., in Lamarck, Encycl. 4: Fevillea cordifolia [var.] hederacea (Poir.) Cogn., in Alph. de Candolle & C. de Candolle, Monogr. Phan. 3: TYPE: On la cultive au jardin des plantes. Elle est originaire de 1 Amerique (v.s.) (HOLOTYPE: P?, n.v.). Fevillea javilla Kunth, in Humboldt, Bonpland, & Kunth, Nov. Gen. Sp. 2: TYPE: CO- LOMBIA. Bolívar: Turbaco, Humboldt & Bonpland 1403 (HOLOTYPE: P, n.v. [microfiche IDC :III. 7]). Fevillea karstenii Cogn., in Alph. de Candolle & C. de Candolle, Monogr. Phan. 3: Nhandiroba karstenii (Cogn.) Kuntze, Revis. Gen. Pl. 1: TYPE: VENEZUELA. DISTRITO FEDERAL: Capaya, near Caracas, Karsten s.n. (HOLOTYPE: W, n.v.; photo ex W: F, MO). The type sheet contains a mixed collection of F. cordifolia and probably Selysia prunifera (Poepp. & Endl.) Cogn. The material of the latter in the lower left portion of the sheet is excluded. Fevillea triloba Sessé & Mociño, Fl. Mexic. ed TYPE: MEXICO. n.v. Siolmatra mexiae Standl., in J.F. Macbride, Publ. Field Mus. Nat. Hist., Bot. Ser. 13(6): TYPE: PERU. LORETO: left bank of Río Marañón, above Rancho Indiana, 110 m, 22 Jan 1932, Mexia 6406 (HOLOTYPE: F; photo ex F: F; ISOTYPES: BM, K, MO, NY, U, US). Fevillea uncipetala Kuhlm., Arch. Jard. Bot. Rio de Janeiro 4: TYPE: BRAZIL. PARÁ: Rio Branco de Obidos, Castanhal Grande, 4 Nov 1919, Ducke s.n. (RB 15924) (HOLOTYPE: RB, n.v.; ISOTYPE: B, destroyed; photograph ex B: F, MO, NY, US). Vine or liana; stem glabrous or lightly to densely pubescent or tomentose; tendrils glabrous to lightly glandular-pubescent. Leaves with the blade unlobed or occasionally 3- to 5-lobed, suborbicular to cordate, (4 )7.5 13( 18) cm long, (2.5 )5 12( 19) cm wide, membranaceous or coriaceous, 5-nerved, the apex acute, the base cordate to truncate or rarely rounded, the main lateral veins terminating in small irregularly shaped glands, the upper and lower surfaces glabrous to densely pubescent, the petiole (2 )3 7 cm long, glabrous or pubescent. Staminate flowers in a paniculate inflorescence on reduced subterminal, lateral branches, the branches subtended by a glandular bract mm long;

10 1980 BRIT.ORG/SIDA 21(4) FIG. 2. Fevillea cordifolia. A. Habit, staminate inflorescence (Lent 3288). B. Staminate flower (Klug 3090). C. Fruit (Huashikat 1078). D. Seed (Huashikat 1078). pedicel 2 4 mm long, glabrous to densely pubescent; bracts linear, glandular, ca. 0.5 mm long; hypanthium 3 4 mm long, glabrous to densely pubescent; calyx shallowly cupular, the lobes mm long, mm wide, obtuse or rounded, densely to sparsely glandular-pubescent, the margin entire, the apex rounded, with a small glandular calycine squamella protruding from each sepal at or near the point of fusion with the petal; petals suborbicular, mm long, mm wide, white, whitish green, cream-colored, light brown, pink or pinkish orange, dark red, or reddish purple, the margin undulate, the median, adaxial ridge with an uncinate appendage; stamens ca. 1 mm long, the anthers ca. 0.5 mm wide, slightly longer than wide, the filaments ca. 0.5 mm long. Pistillate flowers solitary or in pairs; calyx deeply cupular, the lobes suborbicular,

11 ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1981 ca. 3 mm long and wide, pustulate, fleshy; petals oblong, ca. 5 mm long, ca. 3.5 mm wide, cream-colored, the median adaxial appendage slightly raised, with 2 small suborbicular glands on each side of the ridge at the base. Fruit subglobose, gourd-like, cm in diameter, cm long, the surface pustulate, mottled green, zonate above the middle with the hypanthium lip scar and marked at the apex by a raised triradiate line; seeds orbicular, compressed, cm thick, 4 6( 7.5) cm wide, the lateral surface of the woody layer striate-verrucose or pustulate, with the outer edges smooth or occasionally tuberculate, the tubercles 2 3 mm long. Distribution and ecology. Southern Mexico east to Puerto Rico, south through Central America, and into South America to Bolivia. A canopy plant of wet forests from sea level to 1,700 meters. Selected specimens examined. MEXICO. Guerrero: Acapulco, , Palmer 335 (MO). GUATE- MALA. Sacatepéquez: near Antigua, m (cultivated, found in market), Nov 1938 Feb 1939, Standley (F). NICARAGUA. Río San Juan: 1 km E of the village of Sábalo, 11 02'N, 84 29'W, 50 m, 6 Sep 1985, Moreno (MO); between Pueblo de San Juan del Norte Nuevo and La Casa de Ramón Castillo Viajando by San Juanillo, 10 55'N, 83 49'W, m, 7 Jul 1994, Rueda et al (MO); Río Pigibaye, 18 Feb 1995, Rueda et al (MO); Reserva Indio-Maiz, along Río Indio, 11 06'N, 83 58'W, 5 20 m, 19 Sep 1998, Rueda et al (MO). Rivas: Isla Ometepe, Volcán Maderas, Hacienda La Argentina, 11 27' 28'N, 85 31'W, m, 15 Jun 1984, Robleto 864 (MO); Las Cuchillas, Isla Ometepe-Volcán Maderas, 11 27'N, 85 28'W, m, 2 Jun 1985, Robleto 1972 (MO). Zelaya: Caño Monte Cristo, La Grupera, 11 33'N, 87 48'W, ca. 10 m, 4 Feb 1982, Moreno & Sandino (MO, NY). COSTA RICA. Heredia: Finca La Selva, OTS field station on the Río Puerto Viejo just E of its junction with the Río Sarapiquí, 24 Mar 1980, Hammel 8251 (MO). Limón: Tortuguero-Sierpe basin, near Río Sierpe and Río Penetencia, 10 32'40"N, 83 32'50"W, 20 m, 21 Jan 1997, Hammel & Grayum (MO); Río Jiménez, 18 Mar 1973, Lent 3288 (F, MO); Cordillera de Talamanca, Reserva Biológica Hitoy Cerere, road between Estación de la Reserva and Cerere, 9 40'20"N, 83 01'35"W, 100 m, 23 Feb 1989, Herrera & Chacón 2434 (MO). Puntarenas: Península de Osa, Estación de Oro, along the Aquaduct, 08 42'00"N, 83 29'10"W, 150 m, 10 Feb 1996, Angulo 517 (MO, NY); Península de Osa, La Palma, Guadalupe, Finca de Efraín González, 08 38'30"N, 83 28'00"W, 50 m, 17 Aug 1993, Aquilar 2119 (MO); Península de Osa, Rancho Quemado, road to Draque, 08 42'00"N, 83 33'00W, 100 m, 30 Jan 1991, Nielsen 895 (MO); Valle de Coto Colorado, 08 46'00"N, 83 15'00"W, 100 m, 25 Jun 1993, Quesada & Segura 705 (MO); Playa San Josecito, Península de Osa, 08 37'00"N, 83 44'00"W, m, 10 Dec 1993, Quesada et al. 848 (MO); Valle de Coto Colorado, shore of Río Esquinas, mouth of Río Esquinas, 08 44'00"N, 83 20'00"W, 30 m, 17 Dec 1993, Segura et al. 255 (NY); Forest de Santo Domingo de Golfo Dulce, Mar 1896, Tonduz (BR). San José: Cordillera de Talamanca, Las Nubes, Estación Santa Elena, 09 23'30"N, 83 36'30"W, 1150 m, 14 Feb 1996, Alfaro 477 (MO). PANAMA. Chiriquí: Burica Peninsula, Rabo de Puerco, 8 km along road W from Puerto Armuelles, 150 m, 19 Feb 1973, Busey 440 (F, MO, NY, USF). Colón: Barro Colorado Island, 100 m S of Zetek Trail, 600 m, 26 Aug 1970, Croat (F, MO, NY, USF). Darién: Río Sabana, above Sante Fe, 14 Sep 1967, Duke (MO). Los Santos: 17.8 mi S of Macaracas, ca. 300 m, 25 May 1967, Burch 1605 (MO). Panamá: 12.4 km E of Canita, 10 Oct 1975, Witherspoon 8704 (MO). CUBA. Oriente: Bayate at Río Jagua, 4 May 1919, Ekman 9613 (G, K, NY, US). JAMAICA. Portland: gorge of the Swift River at Eden, 0.5 mi N of Paradise, ca. 30 m, 19 Mar 1956, Proctor (MO). St. Andrew: Hope Gardens, 29 May 1902, Harris 8381 (BM, NY). St. Ann: cave near St. Ann s Bay, Dec 1873, Purdie s.n. (K). St. Mary: Wapping Stairs, N side of Guys Hill, 20 Mar 1960, Proctor (NY). St. Thomas: near Dove Hall, 10 Feb 1850, Alexander s.n. (K, NY). HAITI. Massif du Nord, Le Borgne, edge of RiviPre du Borgne, 12 Sep 1925, Ekman 4851 (US). DOMINICAN REPUBLIC.

12 1982 BRIT.ORG/SIDA 21(4) El Seibo: Cordillera Oriental, ca. 6 8 km S of Miches-Las Lgunas de Nisibon Highway, on road to Batey Arroyo Santiago, basin of Río Yeguada (S of Miches), 18 55'N, 69 04'W, m, 28 Jun 1990, Zanoni & Jiménez (MO). La Vega: Jarabocoa, Monabao, Los Calabazos, Arroyo Frio entrance, 19 4'23"N, 70 43'34.6"W, ca. 774 m, 19 Mar 2001, Ososki & Saborío 299 (NY). PUERTO RICO. Along road between Utuado and Adjuntas, km 40, 2 Feb 1997, Acevedo & Angell 9419 (NY); Bayamón, 31 Mar 1885, Sintenis 986 (BM, BR, G, K, M, NY, US). COLOMBIA. Amazonas: Loreto-Yacú River, ca. 100 m, Sep 1946, Schultes & Black (K). Antioquia: near Río León ca km upstream and S of the river mouth ca. 15 km W of Chigorodó, ca. 7 45'N, 76 50'W, ca. 100 m, 14 Mar 1962, Feddema 1907 (NY) ; Murrí la Blanquita, Río Murrí, 06 35'N, 76 50'W, 960 m, 28 Feb 1992, Gentry et al (MO); km 28.8, Nutibara-La Blanquita road, 06 40'N, 76 27'W, 1020 m, 5 Nov 1988, Zarucchi et al (MO). Atlántico: Barranquilla and vicinity, Jan 1934, Elias 1173 (F, US). Bolívar: vicinity of Turbaco, Nov 1920, Heriberto 469 (F, US). Chocó: Río San Juan, Quebrada del Taparal, 5 20 m, 30 May 1946, Cuatrecasas (F); Río Chintado, 1 2 1/2 hrs. above La Nueva, 6 Feb 1967, Duke 9865 (NY); right bank of Río Baudó, ca km upstream of estuary, between estuary of Quebrada Porquera and the sawmill Porquera, ca. 5 m, 6 Feb 1967, Fuchs & Zanella (NY). Cundinamarca: Sierra de Subía, 6.6 km N of Cumaca along road to Viota, 1700 m, 22 Jun 1972, Barclay et al (US). Magdalena: Santa Marta, 5 mi S of Ciénaga, near sea level, 12 Sep , Smith 1607 (BM, BR, F, G, MO, NY, US). Meta: Serranía de la Macarena, Plaza Bonita, bank of Río Güejar, 400 m, 14 Nov 1949, Philipson et al (F, BM, US). Putamayo: Frontera Colombia-Ecuatoriana, along Río San Miguel, at mouth of the Río Conejo, 300 m, 9 Dec 1940, Cuatrecasas (US); San Antonio, Alto Campucana path, Finca La mariposa, m, 01 12'N, 76 38'W, 10 Apr 1 May 1994, Fernández et al (NY). Vaupés: Río Guayabero, 240 m, 8 Nov 1939, Cuatrecasas 7499 (US). VENEZUELA. Apure: Reserva Forestal San Camilo, vicinity of Chirocoa, 9 10 km E of Caserío San Camilo (El Nula), 200 m, 1 Apr 1968, Steyermark et al (MO, NY). Aragua: Parque Nacional Henri Pittier, Estación Biológica de Rancho Grande, 6 Jun 1987, Rojas & Rojas 3617 (MO, NY); Parque Nacional Henry Pittier, between trail up Periquito and Fila de Periquito, along upper slopes of tributary to Quebrada Palo Vaco on side towards Lago Valencia, opposite Rancho Grande Biological Station, m, 25 Oct 1961, Steyermark (F, NY, US). Carabobo: Río Morón, E of Morón, 'N, 'W, 0 50 m, Jul 1991, Diaz & Jiménez 487 (MO). Delta Amacuro: between La Margarita and Puerta Miranda, Río Cure, m, 26 Nov 1960, Steyermark (NY, US); Río Acure, 29 Jan 1980, Trujillo & Sulbaran (MO). Distrito Federal: Hacienda Chichiriviche, ca. 300 m, Jul 1958, Aristeguieta 3225 (NY, US). Mérida: km above dam site on Río Guaimaral, 7 45'N, 71 29'W, 15 Mar 1981, Liesner & González (MO). Miranda: Carretera Santa Teresa Guatopo, ca. 300 m, 4 Jun 1959, Trujillo 4149 (US). Yaracuy: Sierra de Aroa, 2 Jul 1953, Aristeguieta & Pannier 1854 (NY). Zulia: ca. 5 km SSE of Destacamento Guasare No. 1 (La Yolanda), 10 52'10"N, 72 29'30"W, m, 16 Nov 1982, Bunting et al (NY); 8 km from San José de Los Altos on road to Caño Colorado, 390 m, 27 Nov 1977, Jeffrey & Trujillo 2396 (F, K, MO); 6 km W of main road and 2 km S of Río Catatumbo, 09 6'N, 72 42'W, ca m, 29 Mar 1982, Liesner & González 13347(MO, NY). ECUADOR. Morona Santiago: El Centro Shuar Pampants, Río Kankaim (Cangaime), 02 47'S, 77 36'W, 300 m, 10 Sep 1985, Warush RBAE85 (NY). Napo: 5 km SE of Las Sachas, 300 m, 13 Apr 1985, Baker et al (NY); Parroquia Dureno, indigenous Cofán-Dureno community, 00 02'S, 76 42'W, 350 m, Dec 1987, Cerón & Cerón 3107 (MO, NY); Estación Biológica Jatun Sacha, Río Napo, 8 km E of Misahuallí, 01 04'S, 77 36'W, 450 m, 22 Oct 1988, Cerón & Iguago 5510 (MO); Hollín-Loreto-Coca highway, between Avila and Río Pucuno, 00 39'S, 77 22'W, 800 m, 10 Dec 1987, Cerón et al (MO, NY); Parque Nacional Yasuní, along Maxus road and pipeline construction project, km 54 54, Sep 1993, Dik 436 (MO, USF); km 2, new Cotundo- Coca highway, 1130 m, 5 Aug 1984, Dodson et al (MO); Coca-Auca oilfields road, km 53, 00 50'S, 76 52'W, 400 m, 20 Aug 1979, Jaramillo & Coello (NY, US); Maxus road, km 1.8, 00 27'S, 76 38'W, 21 Sep 1997, Klitgaard et al. 617 (NY); Parque Nacional Yasuní, Añangu, along Río Añangu near junction with Río Napo, 0 31'S, 76 23'W, ca. 270 m, 16 Jun 1982, Luteyn et al (NY); near NW corner of Lake Limóncocha, Sep 1969, Mowbray (MO); along Río Indillama, tributary of Río Napo,

13 ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1983 Comuna Pompeya, 00 30'S, 76 40'W, 220 m, 5 Dec 1992, Neill et al (MO, NY); right bank of Río Napo, 8 km from Puerto Misahuallí, 01 04'S, 77 37'W, 450 m, 7 16 Sep 1988, Palacios 3006 (MO, NY); Codo Sinclair, 00 08'S, 77 27'W, 650 m, Sep 1990, Palacios 5719 (MO); Río Huataraco, towards Ishpano, 00 44'S, 77 23'W, 700 m, 30 Nov 1992, Palacios (MO, USF); 6 km N of Shushufindi, towards Dureno, 0 10'S, 76 40'W, 450 m, 23 Sep 1985, Palacios et al. 829 (MO); Dureno on Río Aguarico, 2 Jul 1966, Pinkley 103 (MO); Yusuní National Park, along Maxus road and pipeline construction project, km 21, 00 33'S, 76 31'W, 250 m, 24 Jul 1994, Pitman 646 (MO, USF); 20 km N of Coca, Palmoriente property, 00 20'S, 77 05'W, 250 m, 3 21 Nov 1989, Rubio 332 (MO, NY). Pastaza: ARCO oil well Villano 2, 01 25'S, 77 20'W, 400 m, 1 18 Dec 1991, Hurtado 2908 (MO); Petro-Cañada highway under construction, Via Auca, 115 km S of Coca, 5 km S of Río Tigüino, 01 15'S, 76 55'W, 320 m, 1 6 Mar 1989, Zak 4135 (MO). Pichincha: Santo Domingo de los Colorados, 800 m, 10 Aug 1945, Solís (F). Carchi: Reserva Indígena Awá, Parroquia Tobar Donoso, sector El Baboso, 78 20'W, 00 53'N, 1600 m, 3 Oct 1991, Rubio & Talcuz 295 (MO, USF). PERU. Amazonas: S of Huampami across Río Cenepa, m, 27 Dec 1972, Berlin 716 (MO); Río Santiago, Cantón de la Quebrada Caterpiza, 1 km from community of Caterpiza, 200 m, 11 Sep 1979, Huashicat 552 (MO, USF); Yamayakat Brosque, 04 55'S, 78 19'W, 320 m, 16 Jan 1996, Jaramillo et al. 881 (MO). Huánuco: vicinity of Tingo María Insupte, 670 m, 16 Aug 1961, Schunke 5645 (F, US). Loreto: Flor de Yarina-Río Samiria, 20 Oct 1982, Ayala et al (NY); Explorer s Inn, Río Amazonas near Indiana, 03 30'S, 73 03'W, 130 m, 20 Feb 1988, Gentry et al (MO); Santa Rosa, lower Río Huallaga below Yurimaguas, ca. 135 m, 1 5 Sep 1929, Killip & Smith (NY); Balsapuerto, ca. 220 m, May 1933, Klug 3090 (BM, F, G, MO, NY, US); Quebrada de Tamishaco above Tamishaco, 7 Nov 1978, Rimachi 4046 (MO); Florída, 8 Feb 1980, Rimachi 4856 (NY); Río Itaya, highway to San Antonio, near Venezia, 90 m, 21 Sep 1994, Rimachi (NY); Bosque Nacional de Iparía, along the Río Ucayali near Iparía (80 km at the confluence with Río Pachitea), m, 23 Aug 1968, Schunke 2670 (NY); San Antonio, Río Itaya, 04 10'S, 73 20'W, 150 m, 13 Dec 1982, Vásquez & Jaramillo3597 (MO); Cocha Pastor, Isla Padre, 03 45'S, 76 10'W, 116 m, 21 Dec 1982, Vásquez et. al (NY); Indiana, Explorama Inn, 03 30'S, 73 05'W, 108 m, 12 Apr 1992, Vásquez et al (MO). Madre de Dios: Parque Nacional del Manu, Cocha Cashu Biological Station, 21 Aug 1976, Foster & Augspurger 3394 (K, MO, NY, US); Cocha Cashu Camp, Parque Nacional de Manu, along Río Manu, 380 m, 22 Oct 1979, Gentry et al (NY); Cuzco Amazónico Lodge, 15 km NE of Puerto Maldonado, 12 35'S, 69 03'W, 200 m, 18 Jun 1990, Núñez (MO); Las Piedras, Cusco Amazónico, Río Madre de Dios, 12 29'S, 69 03'W, 200 m, 13 Aug 1991, Timaná 1997 (MO). San Martín: Valley of San Martín, E of Tarapoto, Funde de San Isidro near Codo Creek, 1000 m, 15 Aug 1937, Belshaw 3230 (NY); Pongo de Cainarachi, Río Cainarachi, tributary of Río Huallaga, ca. 230 m, Sep Oct 1932, Klug 2749 (BM, F, G, MO, NY, US); Rioja-Pomacochas road, below Venceremos, ca. 20 km NW of Rioja, 05 45'S, 77 38'W, 1600 m, 8 Feb 1984, Gentry & Smith (MO); above Chazuta, W of Quebrada Chazuta, 06 34'S, 76 12'W, m, 28 Aug 1986, Knapp 8177 (NY); W of Nueva Aspusana (2 hrs down Río Huallaga from La Roca), 8 Aug 1962, Mathias & Taylor 6116 (MO); Fundo La Campiña, 2 km below Tocache Nuevo, right bank of Río Huallaga, ca. 400 m, 23 Aug 1969, Schunke 3377 (NY); Tananta (left bank of Río Huallaga), 6 Oct 1970, Schunke 4479 (NY); Nueva Unión below Puerto Huicte (right bank of Río Huallaga), m, 1 Aug 1974, Schunke 7965 (NY). Ucayali: Bosque Nacional de Iparía, along the Río Ucayali near the village of Iparía (ca. 80 km above the confluence with Río Pachitea), m, 23 Aug 1968, Schunke 2670 (F, G); Río Novia, right bank at native community San José, 10 12'S, 70 57'W, 189 m, 26 Feb 2002, Schunke & Graham S14909 (NY). BRAZIL. Acre: margin of Rio Azul, ca 'S, 73 39'W, 13 Oct 1986, Campbell et al (NY); Sena Madureira, 28 Sep 1980, Cid & Nelson 2596 (NY); basin of Rio Juruá. right bank of Rio Tarauacá, 8 32'51"S, 71 28'39"W, 17 Nov 1995, Daly et al (MO, NY); near mouth of Rio Macauã (tributary of Rio Iaco), 9 20'S, 69 W, 23 Aug 1933, Krukoff 5610 (BM, F, G, M, MO, US). Amazonas: near mouth of Rio Embira (tributary of Rio Tarauacá), 7 30'S, 70 15'W, 6 Jul 1933, Krukoff 5209 (BM, F, G, M, MO, NY, US). Pará: Belém, 20 Dec 1950, Black (NY); Rio Pacajá 2 50'S, 50 50'W, 15 Oct 1965, Prance et al (NY); Rio Mocoões, 45 min. below Frances, 00 45'S, 49 41'W, 10 Nov 1987, Prance et al (MO, NY); Altamira, left bank of lower

14 1984 BRIT.ORG/SIDA 21(4) Rio Xingu, 19 Oct 1986, Souza et al. 385 (NY); Travessão do CNEC, between D13 and edge of Rio Xingu, 2 Dec 1986, Souza et al. 635 (NY); Ilha de Marajó, Cuantã, above Anajás, Rio Anajás, 00 57'S, W, 2 Nov 1987, Tavares 334 (NY). Rondônia: E bank of Rio Madeira at Misericórdia between Cachoeiras Misericórdia and Madeira, 30 Jul 1968, Prance et al (NY). BOLIVIA. Cochabamba: Proyecto Valle del Sacta, km 240 on Santa Cruz-Villa Tunari highway, 17 00'S, 64 46'W, 290 m, Jul 1989, Smith et al (MO). El Beni: Río Beni, above confluence with Río Quiquibey, 3.5 hrs. upstream from Rurrenabaque, 14 44'S, 67 25'W. 320 m, 23 May 1990, Daly et al (MO, NY). La Paz: Parque Nacional Madidi, 10.2 km NW of turnoff in Tumupasa, km from summit, 14 09'57"S, 67 55'02"W, 830 m, 9 Aug 2000, Croat et al (MO); Santa Fe, NE of community,13 40'S, 68 12'W, 250 m, 10 Aug 1995, DeWalt et al. 823 (MO, NY); basin of Río Bopi, San Bartolomé near Calisaya, m, 1 22 Jul 1939, Krukoff (F, G, K, MO, NY, US); Parque Nacional Madidi, near Arroyo Aguapolo and Río Tuichi, 270 m, 16 Mar 2002, Macía et al (NY). Santa Cruz: Parque Nacional Amboró, along Río Isamá [ Río Pitasamá ], 'S, 'W, 475 m, 11 Oct 1990, Nee (NY); Estancia San Rafael de Amboró, 15 km (by air) SSE of Buena Vista, 17 35'S, 63 37'W, 375 m, 28 Jul 1987, Nee et al (NY); Río Palometilla, 400 m, 16 Jun 1927, Steinbach 7904 (F, G, MO, NY, BM). Fevillea cordifolia is a polymorphic species exhibiting considerable variation throughout its range. The species can be readily distinguished from other members of the genus by its conspicuous marginal laminar glands. It is most similar to F. trilobata of eastern Brazil and is distinguished from that species by the typically rounded leaf lobes (when lobes are present), rather than the angular lobes characteristic of F. trilobata. Fevillea trilobata also has a distinctive stipitate-glandular pubescence on the hypanthium of the staminate flowers, which is sharply differentiated from the uncinate pubescence of pedicel. This pattern of pubescence is lacking in F. cordifolia. Fevillea moorei Hook. f., Bot. Mag. t Nhandiroba moorei (Hook. f.) Kuntze, Revis. Gen. Pl. 1: TYPE: GUYANA or BRAZIL: Cultivated at Liverpool Botanic Garden, 1871, Ty reman s.n. (HOLOTYPE: K; ISOTYPE: BR-fragment). Vine or liana; stem glabrous; tendrils glabrous. Leaves with the blade unlobed, broadly ovate, 6 12 cm long, cm wide, membranaceous, drying light green, 3-nerved, the apex acuminate, the base rounded, with 2 small, irregularly shaped glands near the petiole, the upper and lower surfaces glabrous, the petiole cm long, glabrous. Staminate flowers in a racemose inflorescence ca. 6 cm long, (10 12 cm fide Hooker f.); pedicel 1 4 mm long, glabrous; hypanthium 5 6 mm long, glabrous; bracts linear, ca. 0.5 mm long; calyx shallowly cupular, the lobes oblong, ca. 6 mm long, 4 mm wide, the apex obtuse, the margin entire, with a glandular calycine squamella protruding at or near the point of fusion with the petals; petals suborbicular, somewhat broader at the apex, ca. 1 cm long, 1 cm wide, pale brick-red, the margin undulate; stamens ca. 4 mm long, the anthers ca. 1 mm long, slightly longer than wide, the filament ca. 3 mm long. Pistillate flowers and fruit not seen. Distribution and ecology. Known only from the type material received by J.D. Hooker from David Moore, curator of the Glasnevin Botanic Garden, Dublin, Ireland, who received it from Mr. Tyreman of the Liverpool Botanic Garden.

15 ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1985 The material was labeled as Strychnos curari and said to have come from Gambia. As suggested by Hooker in the protologue, it is probable that the plant is from South America, possibly Guyana or the Amazon River basin of Brazil where Strychnos toxifera, the source of the drug curare is native and was known to occur at that time. Hooker named the plant in honor of his friend, Dr. Moore. Fevillea moorei is most similar to F. bahiensis of southern Bahia, Brazil. It is easily distinguished by its larger staminate flowers, its leaf margins lacking uncinate trichomes, and its 3-nerved leaves which dry to a light green color. Fevillea pedatifolia (Cogn.) C. Jeffrey, Kew Bull. 16: (Fig. 3). Alsomitra pedatifolia Cogn., in Martius, Fl. Bras. 6(4): Siolmatra pedatifolia (Cogn.) Cogn., in Engler, Pflanzenr. 4(Heft 66): TYPE: BRAZIL. AMAZONAS: Rio Solimões, , Martius s.n. (HOLOTYPE: M; photo ex M: F, MO, NY, US; ISOTYPES: BR-fragment). Alsomitra peruviana Huber, Bol. Mus. Paraense Hist. Nat. 4: Siolmatra peruviana (Huber) Cogn., in Engler, Pflanzenr. 4(Heft 66): Fevillea peruviana (Huber) C. Jeffrey, Kew Bull. 16: TYPE: PERU. UCAYALI: Río Ucayali, near Canchahuaya, 28 Oct 1898, Huber 1390 (HOLOTYPE: MG, n.v.; photo ex MG: F; ISOTYPE: F-fragment). Siolmatra amazonica Cogn., in Engler, Pflanzenr. 4(Heft 66): Fevillea amazonica (Cogn.) C. Jeffrey, Kew Bull. 16: TYPE: BRAZIL. AMAZONAS: Rio Juruá, Juruá Miry, Oct 1901, Ule 5819 (LECTOTYPE: here designated, K; ISOLECTOTYPES: F-fragment, MG, n.v.; photo ex MG: F; B, destroyed; photo ex B: F, MO, NY, US). Siolmatra simplicifolia Harms, Notizbl. Bot Gart. Berlin-Dahlem 11: Fevillea simplicifolia (Harms) C. Jeffrey, Kew Bull. 16: TYPE: PERU. LORETO: Lower Río Huallaga, Yurimaguas, Puerto Arturo, 15 Nov 1929, Williams 5072 (HOLOTYPE: B, destroyed; LECTOTYPE: here designated, F; ISOLECTOTYPE: US). Vine or liana; stem glabrous to densely glandular-pubescent; tendrils sparsely pubescent or glabrous. Leaves with the blade unlobed, or 3- to 5-lobed, or 3- to 5-foliolate, membranaceous to coriaceous, the unlobed or the lobed leaves ovate to ovate-oblong, (6 )8 15( 18) cm long, 9 14( 17.5) cm wide, 5- to 7-nerved, the divisions of the lobed leaves or the leaflets of the foliolate leaves (6 )8 15( 18) cm long, 4 8 cm wide, 1- to 2-nerved, with a petiolule up to 2 cm long, the apex of the blade or leaf divisions acuminate, the base of the blade or leaf divisions oblique or rounded, the margin entire or coarsely crenate-toothed, the primary lateral veins terminating in a small irregularly shaped marginal gland, the upper and lower surfaces glabrous or lightly pubescent, especially along the leaf veins, the petiole 3.5 5( 7) cm long, glabrous to sparsely pubescent, with 2 opposite, prominent, irregularly shaped median to subapical glands. Staminate flowers in a paniculate inflorescence on reduced lateral branches, each branch subtended by a thin scale-like glandular-pubescent bract mm long; pedicel mm long, sparsely glandular-pubescent to glabrate; hypanthium mm long, sparsely glandular-pubescent to glabrate; calyx shallowly cupular, the lobes light green or greenish brown, suborbicular, mm long and wide, the apex rounded or obtuse, glandular-pubescent or glabrous, the margin slightly erose and glandular-ciliate, with a glandular calycine squamella

16 1986 BRIT.ORG/SIDA 21(4) FIG. 3. Fevillea pedatifolia. A. Habit, staminate inflorescence (Klug 4187). B. Staminate flower (Klug 2678). C. Fruit (Barbour 5533). D. Seed (Barbour 5533). protruding at or near the point of fusion with the petals; petals suborbicular, 1 2 mm long, mm wide, white or greenish white, greenish yellow, or yellowish, the margin slightly erose, the median adaxial ridge slightly raised; stamens mm long, the anthers ca mm long, slightly longer than wide, the filament ca. 0.5 mm long. Pistillate flowers in pairs; pedicel mm long; calyx deeply cupular, the lobes ca. 3 mm long, ca. 2 mm wide, dark greenish brown, the surface pustulate; petals white. Fruit subglobose, ca. 14 cm long, 13 cm wide, gourd-like, indehiscent or rarely circumscissile dehiscent along the hypanthium lip scar (A. Gentry, pers, comm.), the surface smooth or pustulate, usually a mottled green color; seeds orbicular, compressed, 4 5 cm long and

17 ROBINSON AND WUNDERLIN, REVISION OF FEVILLEA 1987 wide, cm thick, silver-white upon drying, the woody layer smooth to slightly pustulate, the winged margin ca. 0.5 cm wide. Distribution and ecology. Amazonian Ecuador and Peru, south to Bolivia, and east to Acre and Amazonas, Brazil. A plant of moist to wet forests, occurring at meters in elevation. Selected specimens examined. ECUADOR. Napo: Estación Biológica Jatun Sacha, 8 km E of Misahuallí, 01 04'S, 77 36'W, 400 m Jan 1989, Cerón 6076 (MO); km 2, new road from Cotundo to Coca, 1130 m, 5 Aug 1984, Dodson et al (NY). PERU. Amazonas: 1 km from La Poza, W of Río Santiago, 180 m, 21 Aug 1979, Huashikat 111 (MO); valley of Río Santiago, Quebrada Caterpiza, 2 3 km behind the community of Caterpiza, 3 50'S, 77 40'W, 200 m, 8 Feb 1980, Tunqui 783 (MO). Ayachucho: between Santa Rosa and Hacienda Luisiana, 640 m, 9 Sep 1976, Wasshausen & Encarnación 632 (NY). Huánuco: Codo de Puzuzo, floodplain of Río Puzuzo, S of settlement to main river, 9 40'S, 75 25'W, 450 m, 21 Oct 1982, Foster 9370 (USF). Junín: Río Negro, 800 m, 14 Aug 1960, Woytkowski 5795 (G, MO, US). Loreto: Yanamono, Explorama Tourist Camp on Río Amazonas between Indiana and mouth of Río Napo, 03 28'S, 72 48'W, 120 m, 26 Jul 1980, Gentry et al (MO); Yanamono, Explorama Tourist Camp, Río Amazonas halfway between Indiana and mouth of Río Napo, 03 28'S, 72 50'W, 130 m, 13 Jul 1983, Gentry et al A (MO); Balsapuerto, 220 m, Feb 1933, Klug 2896 (BM, F, G, MO, NY, US); Indiana, Reserva Explorama (Yanamono), 03 30'S, 72 50'W, 90 m, 28 Sep 1990, Pipoly et al (MO); Yanamono tourist camp, 50 mi. NE of Iquitos, 3 30'S, 72 50', ca. 106 m, 19 Oct 1980, Vásquez & Jaramillo 586 (NY); Indiana, Iquique, 03 30'S, 72 58'W, 115 m, 16 Dec 1987, Vásquez & Jaramillo (MO); Indiana, Explorama Reserve, 03 28'S, 72 50'W, 106 m, 9 Nov 1989, Vásquez & Jaramillo (MO). Madre de Dios: Tambopata, 12 50'S, 69 17'W, 260 m, 8 Nov 1988, Alexiades et al. 81 (MO, NY); Río Piedras, near confluence with Río Pariamanu, 12 40'S, 69 17'W, 260 m, 11 Jan 1991, Alexiades et al (NY); Lago Tres Chimbadas, ca river km SSW of Puerto Maldonado, ca air km NW effluence of Río La Torre (Río D Orbigny) Río Tambopata, 12 49'S, 69 17'W, ca. 260 m, 7 Jun 1980, Barbour 5533 (MO); Río Manu, Cocha Cashu Biologial Station, 350 m, 1 Oct 1980, Foster 5439 (NY); Cocha Cashu, vicinity of ox-bow lake of Río Manu, between Panaqua and Tayakome, Aug 1974, Foster et al (K, MO, US). Las Piedras, Cusco Amazónico, 12 29'S, 69 03'W, 200 m, 24 Jun 1991, Timaná 1844 (MO). Pasco: Río Pichis, 1 hr. below Puerto Bermudez, between Puerto Bermedez and Paujil, 10 10'S, 74 50'W, 200 m, 17 Jun 1983, Gentry et al (MO); Iscozacín, near confluence of Río Palcazu and Río Iscozacín, 10 12'S, 75 13'W, 430 m, 16 Jun 1982, Smith 1911 (NY); Palcazu Valley, Río Chuchurras drainage, 10 09'S, 75 20'W, 400 m, 13 May 1983, Smith 4009 (MO, US). San Martín: Pongo de Cainarachi, Río Cainarachi, tributary of Río Huallaga, 230 m, Sep-Oct 1932, Klug 2755 (BM, F, G, K, MO, NY, US); Quebrada Cachiyacu de Huaquisha, ca m, 7 Dec 1980, Schunke (NY). Ucayali: Bosque Nacional von Humboldt, Pucallpa-Tingo María Road, 8 40'S, 75 0'W, 250 m, 13 Feb 1981, Gentry et al (MO, USF). BRAZIL. Acre: N bank of Rio Juruá opposite Cruzeiro do Sul, 27 Oct 1966, Prance et al (K, MO, NY, US). Amazonas: near Palmares, 11 Sep 26 Oct 1936, Krukoff 8475 (BM, BR, F, G, K, MO, NY, US). BOLIVIA. La Paz: basin of Río Bopi, Asunta (near Evenay), m, Jul 1939, Krukoff (F, G, K, MO, NY). Plants with 3- to 7-foliolate leaves are easily distinguished from other species of Fevillea. They previously had been considered a species of Siolmatra because of this feature, but the presence of the foliar glands, the bilocular anthers, and the large globose fruit with large unwinged seeds clearly separates it. Siolmatra, in contrast, lacks foliar glands, has unilocular anthers, and has a cylindric fruit with winged seeds. Fevillea peruviana and F. amazonica were separated from F. pedatifolia on the basis of leaflet number (3 rather than 5). Fevillea amazonica was further